Flavor

Elizabeth A. Baldwin
USDA/ARS, Citrus and Subtropical Products Laboratory, Winter Haven, FL

Introduction: The quality of fresh produce has traditionally been based on external characteristics of size,
color, and absence of surface defects. Fruit and vegetable breeders select for color, size, disease resistance,
yield and other easily quantified horticultural traits. Because flavor and texture characteristics were not a
part of the selection process, improvements in these quality attributes have not kept pace with the more
easily quantified traits. Public and industry organizations are increasingly concerned with the public’s
growing dissatisfaction over the flavor and texture of some horticultural produce.
Flavor and aroma are perhaps the most elusive and subjective of quality traits. Flavor is taste plus odor
and is mainly composed of sweetness, sourness, and aroma, which corresponds to sugars, acids, and
volatiles. Other components of flavor include bitterness, eg., related to sesquiterpene lactones in chicory
(Peters and Amerongen, 1998), saltiness due to various natural salts, and astringency related to flavonoids,
alkaloids (DeRovira, 1997), tannins (Taylor, 1993), and other factors. The perception of sweetness, ie.,
sugars, one of the most important components of fruit or vegetable flavor, is modified by sourness or acid
levels, and aroma compounds. The contribution of aroma to the flavor quality of fresh produce has gained
increasing attention.
Genetics is the primary determinant of flavor of fresh produce (Baldwin et al., 1991b and 1992;
Cunningham et al., 1985), with pre-harvest environment (Romani et al., 1983), cultural practices (Wright
and Harris, 1985), harvest maturity (Fellman et al., 1993; Maul et al., 1998; Baldwin et al., 1999a), and
postharvest handling (Mattheis et al., 1995; Fellman et al., 1993; Baldwin et al., 1999a and b) having lesser
effect. Fruit such as apples and bananas that continue to ripen after harvest are termed climacteric, while
those such as citrus and strawberries that do not ripen after harvest are termed non-climacteric. The flavor
quality of non-climacteric fruit generally declines after harvest, while climacteric fruit can reach their best
flavor after harvest. Climacteric fruit develop better quality if harvested after the start of ripening, while
fruit of both will be inferior in quality if harvested immature, even if held under optimal postharvest
conditions.
Human perception of flavor is exceedingly complex. Taste is the detection of nonvolatile compounds
(in concentration of parts per hundred) by several types of receptors in the tongue for sugars or
polyalcohols, hydronium ions, sodium ions, glucosides and alkaloids, etc. These correspond to the
perception of sweet, sour, salty and bitter tastes in food. Aroma compounds can be detected in ppb
concentrations and are detected by olfactory nerve endings in the nose (DeRovira, 1997). The brain
processes information from these senses to give an integrated flavor experience. This integration makes it
difficult to determine the relative importance of each input since the brain can interpret changes in aroma
as changes in taste (O’Mahony, 1995) or vice versa. For example, the levels of aroma compounds
influenced panelist perception of sweetness and sourness for tomatoes (Lycopersicon esculentum Mill.)
(Baldwin et al., 1998). Conversely, levels of taste components influenced panelist perception of aromatic
descriptors in mango (Mangifera indica L.) (Malundo et al. 2000a). The perception of certain combinations
of chemicals is synergistic, while others combinations mute our perception in a process called masking. In
contrast to masking, is anesthetization or blanking in which olfactory receptors become overloaded. Lighter
aroma volatiles, eg., top notes, low molecular weight, polar, hydrophilic compounds, are perceived first and
generally have the major impact on perception, while heavier compounds are perceived later, eg.,
background. notes, high molecular weight, non-polar, hydrophobic compounds (DeRovira, 1997).

Sensory Evaluation: Human perception of flavor can be determined by sensory evaluation by taste panels.
Consumer preference and acceptance varies due to socioeconomic, ethnic and geographical background;
often necessitating the segmenting of sub-populations for a particular study (O’Mahony, 1995). Generally
a large number of panelists (ex., 50 to 100) rank their perceptions on a traditional 9-point hedonic scale, but
sometimes a simple 3-point scale including the descriptive terms outstanding, acceptable, and unacceptable
can be effective for tomato fruit evaluation. In one study, adaptation of logistic regression from medical
science proved useful, where a 0 or 1 indicates whether the consumer would or would not purchase a mango
(Mangifera indica). The consumer was asked to base their decision on flavor that was then related to
chemical constituents (Malundo et al., 2000b). Difference testing can be used to measure slight differences
between foods (usually due to one particular aspect of flavor), and is considered a narrow band approach.
Descriptive analysis measures intensities of a set of sensory attributes and is considered a broadband
approach (O’Mahony, 1995). Panelists are trained to detect a range of flavor attributes and score their
intensity, generally on a 150 mm unstructured line. Sensory studies for fresh produce can be used to identify
optimal harvest maturity, evaluate flavor quality in breeding programs, determine optimal storage and
handling conditions, assess effects of disinfestation or preconditioning techniques on flavor quality, and
measure flavor quality over the postharvest life of a product.

Taste Components: Fructose, sucrose, and glucose are the sugars that affect the perception of sweetness
in fruits and vegetables. Fructose is the sweetest, and glucose is less sweet than sucrose. A single “sucrose
equivalent” value is the weighed average of these various sugars in a sample (Koehler and Kays, 1991).
Sugar content is commonly accepted to be synonymous with SSC (soluble solids content), and an
inexpensive refractometer can easily measure SSC. However, the quantification of individual sugars
requires complicated laboratory analysis. Breeders often select for higher SSC in an attempt to increase
sweetness. In some fruits, such as orange (Citrus sinensis), SSC relates to sweetness, while in others, like
tomato and mango, the relationship is not linear (Baldwin et al., 1998 and 1999a; Malundo, et al. 2000a).
Organic acids, such as citrate in citrus and tomatoes, tartaric acid in grapes (Vitis sp.), and malic acid
in apples (Malus pumila), give fruit and vegetables their sour flavor. Some fruits, like melon (Cucumis
melo) or banana (Musa sp.), have very little acid (Wyllie et al., 1995). Different acids can affect sourness
perception depending on their chemical structure. An increase in carboxyl groups decreased acidity, while
an increase in molecular weight or hydrophobicity increased sourness (Hartwig and McDaniel, 1995). For
example, acetate was perceived as more intensely sour than lactic or citrate.
Acids can be measured individually by HPLC (Baldwin et al., 1991a and b), by titration (TA) with
sodium hydroxide (Jones and Scott, 1984), or by pH (Baldwin et al., 1998). Sometimes SSC, the ratio of
SSC/TA, or pH relate better to sourness than TA itself (Baldwin et al., 1998; Malundo et al., 2000a).

Aroma Components: Volatiles that we can perceive contribute to food flavor. The level at which a
compound can be detected by smell (the odor thresholds) can be established in a background similar to a
food medium as described by the Ascending Method of Limits of the American Society for Testing and
Materials (ASTM, 1991). Log odor units are calculated from the ratio of the concentration of a component
in a food to its odor threshold. Compounds with positive odor units contribute to food flavor. Buttery
(1993), for example, determined concentrations, odor thresholds and log odor units for those tomato
volatiles present at levels of one ppb or more (about 30 of > 400 identified compounds). However, the
aroma perception of volatile compounds is affected by the medium of evaluation. For example, both the
thresholds and descriptors of some volatile compounds in tomato were different if the background media
contained levels of methanol and ethanol similar to that found in fresh tomato homogenate or in deodorized
homogenate itself, compared to water (Tandon et al.. 2000) (Table 1).
Aroma compounds are often only released upon cell disruption when previously compartmentalized
enzymes and substrates interact (Buttery, 1993). Some aroma compounds are bound to sugars as glycosides
(celery [Apium graveolens], lettuce [Lactuca sativa]), or glucosinolates (cabbage [Brassica oleracea],
radish [Raphanus sativus]). This linkage can be cleaved by enzyme action or heat during cooking. Others
are breakdown products of lipids, amino acids, lignin, or pigments (Buttery and Ling, 1993).
Measurement of aroma compounds is difficult and time consuming. Earlier studies employed the
classical flavor isolation procedures of steam distillation and/or solvent extraction (Teranishi and Kint,
1993). The disadvantage of this method is that it can qualitatively and quantitatively modify the flavor
profile of a sample (Schamp and Dirinck, 1982). This method is not easily applied to large numbers of
samples, and internal standards must be incorporated to determine recovery. The resulting concentration of
material, however, allows identification of compounds by gas chromatography-mass spectrometry
(GC-MS). More recently, investigators have employed purge and trap headspace sampling methods which
involve trapping and concentrating volatile components on a solid support. Volatiles are later released from
the trap using heat for analysis by GC-MS. This method is excellent for quantification and identification of
aroma compounds (Teranishi and Kint, 1993; Schamp and Dirinck, 1982).
Static headspace methods are said to more closely reflect the true flavor profile, but compounds are
present at low levels, and some may not be detected. Cryofocusing (cold trap) of static headspace volatiles
(Teranishsi and Kint, 1993) reduces this problem since samples are concentrated without heating that may
cause adulteration. This method has been used for quantification of orange juice volatiles (Moshonas and
Shaw, 1997). The newest method is solid phase micro extraction (SPME), a rapid sampling technique
where volatiles interact with a fiber-coated probe inserted into the sample headspace. The probe is then
transferred to a GC injection port where the volatiles are desorbed. It has been used on apples, tomatoes
(Song et al., 1997) and strawberries (Golaszewski et al., 1998; Song et al., 1997).
Aside from GC and GC-MS methods, there are new sensors available that have a broad range of
selectivity. These sensor arrays, called “electronic noses,” are useful to discriminate among samples based
on the interaction of volatile components with the various sensors. The resulting response pattern allows a
particular sample or flavor component(s) to be detected by pattern recognition. However, these instruments
do not give information that leads to identification/quantification of individual compounds. Four basic
sensor technologies have been commercialized to date. Metal oxide semiconductors (MOS), metal oxide
semiconductor field effect transistors (MOSFET), conducting organic polymers (CP), piezoelectric crystals
(bulk acoustic wave; BAW) or quartz crystal microbalance. Such sensors are divided into two classes since
they either operate “hot” (MOS, MOSFET) or cold (CP, BAW). The “hot” sensors are less sensitive to
moisture and have less carry-over from one measurement to the next. The next generation of electronic
noses may use fiberoptic, electrochemical and bi-metal sensors currently under development (Schaller et
al., 1998).

Relating Sensory to Chemical Data: Chemical analysis of flavor compounds provides little insight into
the actual flavor experience. However, sensory attributes, preferences, and decisions can be statistically
related to chemical components in foods (Martens et al., 1994). Correlation of physical measurements with
sensory analysis gives meaning to instrumental data; as was shown with apple and tomato (Baldwin et al.,
1998). For example, linear regression established relationships between levels of sesquiterpene lactones and
bitterness in chicory (Peters and Amerongen, 1998). Multivariate methods require large data sets, but
non-linear regression techniques such as principle component or discriminate analysis yielded useful results
for citrus (Moshonas and Shaw, 1997), strawberry (Fragaria ananassa) (Shamaila et al., 1992), and tomato
(Maul et al., 1998). Differences between samples were found based on measurement of volatiles or other
flavor compounds. Alternatively, sniff ports (olfactometry detectors) can be used with GCs, allowing a
person to determine if odors are detectable as well as their relative intensity as the volatile components are
separated by the GC column. This technique was used on apples (Cunningham et al., 1985; Young et al.,
1996). Descriptive terms can be assigned to the respective peaks on the GC chromatogram that have odor
activity (Acree, 1993). The drawback to this method is that the interactive effects of volatile compounds
with each other and with sugars and acids, both chemically and in terms of human perception, are
eliminated.

Factors that can Affect Flavor of Fruits and Vegetables:

Effect of genetics on flavor
Fruit and vegetable varieties differ in flavor based on sensory and chemical analysis. “Charm” analysis
combines separation on a GC column with a sniff port to assign biological activity (odor activity) to
individual aroma components as they are identified and quantified by GC (Cunningham et al., 1985). This
study with 40 cultivars showed that apple aroma was not the result of the same compounds in every cultivar,
although some common volatile compounds were important in all cultivars.
Important aroma-specific compounds for strawberry included ethyl butanoate, methyl butanoate,
γ-decalactone, and 2-heptanone (Larsen et al., 1992). Strawberry cultivars differed in flavor intensity and
sweetness according to a trained sensory panel (Podoski et al., 1997). Concentrations of several important
compounds including α- and ß-ionones, were higher in wild compared to cultivated raspberries (Rubus sp.).
In addition, numerous aroma compounds were found only in wild berries, all of which may contribute to the
stronger and more pleasant aroma of wild berries (Martin and MacLeod, 1990). In tomato, the TA/SSC
(Stevens et al., 1977) and levels of flavor volatiles varied significantly among varieties (Baldwin et al.,
1991a and b). Insertion of the rin gene to reduce ethylene production and slow tomato fruit softening,
resulted in some deterioration in flavor quality (Baldwin et al., 2000) and reduction in flavor volatiles
(Baldwin et al., 1992; Baldwin et al., 2000). Flavor appears to be related to ethylene production (Baldwin
et al., 1991a and 2000). Transgenic fruit with antisense aminocyclopropanecarboxylic acid (ACC) synthase
(enzyme in the ethylene biosynthetic pathway) had lowered levels of many important flavor volatiles
(Baldwin et al., 2000). Fruit with antisense pectinmethylesterase (demethylates pectin in cell walls) had
lowered levels of methanol, while those with downregulated phytoene synthase (phytoene is a precursor of
carotenoids) had lowered levels of carotenoid-derived volatiles (Baldwin et al., 2000).

Effect of Pre-harvest Factors

Pre-harvest factors such as sunlight, water availability, fertilization, and chemical applications affect crop
growth, and can affect internal quality characteristics of the harvested product, including flavor. Pre-harvest
treatment with aminoethoxyvinylglycine (AVG) suppressed volatile production in pears by 50%, which
was reversed by ethylene exposure (Romani et al., 1983), and heavy rains prior to harvest dilute flavor
compounds in tomato. Fruit from tomato plants treated with increased levels of N and K fertilizer scored
lower in sensory analysis, and showed increased levels of TA, SSC, and several volatiles (Wright and
Harris, 1985). Pre-harvest mite control resulted in sweeter and more flavorful field grown strawberries than
those receiving no treatments, according to a trained sensory panel (Podoski et al., 1997).

Effect of Harvest Maturity

Horticultural crops should be harvested at optimal eating quality, but practical considerations dictate that
they are harvested at a stage that minimizes physical damage during shipping and handling, and maximizes
shelf life. The climacteric stage at harvest affected ester formation in apples (Fellman et al., 1993). Harvest
maturity affected both the sensory and chemical analysis of ripened tomato fruit (Maul et al., 1998).
Tomatoes harvested at the immature green stage resulted in ripened fruit with lower volatile levels than
mature green-harvested tomatoes. Harvest maturity also affected consumer acceptability for mango, and
trained descriptive panel ratings for sweetness, sourness, and various aroma descriptors. Fruit harvested
later were sweeter, less sour and generally had more intense aroma characteristics (Baldwin et al., 1999a).

Effect of Postharvest Handling

Various techniques are used to extend the shelf-life of fruits and vegetables after harvest, to control
postharvest decay, and to eliminate pests (quarantine treatments). These storage techniques and treatments
involve cold, heat, irradiation, chemical applications, and different storage atmospheres.
Tomato fruit stored at 36, 41, 50, and 55 °F (2, 5, 10, and 13 °C) had reduced levels of important
volatiles and had less ripe aroma and flavor as well as more off-flavor compared to fruit stored at 68 °F (20
°C), as quantified by a trained descriptive panel (Maul et al., 2000). Subjection of fruit to heat treatments for
pre-conditioning and decay control (McDonald et al., 1996), resulted in altered aroma volatile profiles. Heat
treatment of apples to reduce physiological and pathological disorders inhibited emission of volatile esters
important to apple flavor (Fallik et al., 1997). Levels of fructose and glucose, but not sucrose, decreased
with increased storage time and storage temperature for muskmelon. However, sensory analysis did not find
differences in flavor or sweetness between stored and freshly harvested melons (Cohen and Hicks, 1986).
CA storage altered flavor of apples, and if prolonged, reduced volatile emission compared to air-stored
fruit, especially lipid-derived esters (Mattheis et al., 1995). Low O2 storage decreased ester content and the
enzymatic activity responsible for ester biosynthesis in apples (Fellman et al., 1993). However, when
atmospheres induced anaerobic metabolism, large concentrations of ethanol and acetaldehyde accumulated.
The altered synthesis of fruit volatiles resulted in increased amounts of ethyl acetate and certain ethyl esters
at the expense of others. Sensory analysis of CA-stored apples revealed that intensity of fruity and floral
descriptors decreased after 10 weeks in CA, while sourness and astringency were higher compared to apples
stored in air. CA storage also increased certain volatiles in tomato, compared to air-stored fruit (Crouzet et
al., 1986).
Use of packaging and edible coatings can create a modified atmosphere (MA) with reduced O2 and
elevated CO2 levels, similar to that of CA. Lowering O2 and raising CO2 can maintain the quality of many
fresh fruits and vegetables for extended periods. However, exposure of fresh produce to O2 levels below
their tolerance level can increase anaerobic respiration and lead to the development of off-flavor. Use of
edible coatings affects flavor and the level of volatile flavor compounds in citrus (Cohen et al., 1990), apple
(Saftner et al., 1999) and mango fruit (Baldwin et al., 1999b). The coating barrier probably induced
anaerobic respiration and the synthesis of ethanol and acetaldehyde, and entrapped volatiles, including
ethanol and acetaldehyde (Baldwin et al., 1999b). In broccoli, sulfur-containing volatiles, including
methanethiol and dimethyl disulfide, are produced in response to anaerobic conditions that can be created
by MAP (Dan et al., 1997). Storing strawberries in MAP altered volatile profiles depending on conditions
(CO2, mixed gases, or air), enabling separation of the samples using multivariate statistics (Shamaila et al.,
1992). Fruit treated with CO2 had the greatest change in volatile levels. This was confirmed by another
study where strawberry fruit stored in a CO2 saturated atmosphere exhibited significant changes in volatile
levels and phenylalanine ammonia lyase (PAL) activity (Dourtoglou et al., 1995). The amino acid
phenylalanine is the precursor to a number of volatiles through a pathway for which PAL is the key enzyme.
In addition to CA, other gaseous treatments of fruits and vegetable have been reported. Use of ethylene
to synchronize ripening has been practiced for years on banana and tomato, and for degreening of citrus.
Ethylene gassing of tomato fruit alters volatile levels (McDonald et al., 1996). Treatment of apple fruit with
1-methylcyclopropene (1-MCP) and methyl jasmonate inhibited both ethylene production and production
of many volatile alcohols and esters, including the formation of esters from alcohols (Fan and Mattheis,
1999). Treatment of bananas with 1-MCP also suppressed volatile production and composition, resulting in
an increase in alcohols and a decrease in related esters (Golding et al., 1999). Application of acetaldehyde
and ethanol vapors to blueberries, tomatoes and pears increased their sugar content, sugar-acid ratio, and
hedonic sensory rating (Paz et al., 1981).
Other chemical treatments of fresh produce may also affect flavor. For example, pressure infiltration
of apples with calcium chloride transiently reduced levels of important flavor volatiles (Saftner et al., 1999).

Flavor of some Popular Fruits and Vegetables:

Apple
Sucrose is the major sugar in apples, although it is slowly hydrolyzed to glucose and fructose during latter
ripening stages. The major organic acid is malate, although some citrate is also present (Knee, 1993).
Eleven aroma compounds contribute to apple aroma in most of the 40 cultivars, while 27 other compounds
contributing to flavor where found only in certain genetic types (Cunningham et al., 1985). Loss of apple
flavor after long term CA storage is a major problem, probably due to the reduction of volatile synthesis
during storage (Mattheis et al.1995).

Peach (Prunus persica)

The main sugar in peaches is sucrose, but cultivars differ greatly in glucose:fructose:sorbitol ratios which
may contribute to differences in flavor. The major organic acids are malate and citrate, with malate levels
declining and citrate levels increasing as fruit ripen (Brady, 1993). Aroma of peaches and nectarines is
distinguished by the presence of γ- and γ-lactones (peach-like and coconut-like, respectively), although
other esters and aldehydes contribute to peach flavor (Do et al., 1969; Crouzet et al., 1990). γ-Lactones from
C-5 to C-12, δ-lactones and unsaturated lactones represent over 25% of the volatiles, with γ-lactone being
the second most abundant component after benzaldehyde. γ-Undecalactone, although rarely reported in
natural extracts, has a distinct peach odor. It has been named ‘peach aldehyde,’ and is used in peach flavor
formulations (Crouzet et al., 1990). Ethyl hexanoate and 3-methylbutanoate, linalool, α-terpineol,
6-pentyl-α-pyrone (coconut odor) and benzyl alcohol are also considered important (Crouzet et al., 1990).

Small Fruits
Strawberry: In most berry fruits sucrose, glucose and fructose are present in roughly equivalent
concentrations (Manning, 1993), and citrate is the major organic acid. Over 200 volatile compounds have
been identified in strawberry. C-6 aldehydes such as hexanal and trans-2-hexenal are found, as well as
lipoxygenase and hydroperoxide lyase. Lipoxygenase acts on linolenic acid to form 13- and
9-hydroperoxides which are cleaved by hydroperoxide lyase to form hexanal and cis-3-hexenal. The cis-3
hexenal is then isomerized to trans-2-hexenal (Perez et al., 1999), as was reported for tomato (Galliard, et
al., 1977; Riley et al., 1996). 2,5-Dimethyl-4-hydroxy-3(2H)-furanone (furaneol) and its methyl ether
(mesifuran) are important aroma components in both strawberry and tomato and are considered to be
glycosidically bound in both fruits (Roscher et al., 1997). Of over 100 volatile compounds identified from
strawberry, furaneol, ethyl hexanoate, and ethyl butanoate are considered to be the character impact
compounds (Zabetakis and Holden, 1997). Sensory analysis of strawberry juice showed that furaneol was
positively related to fresh flavor and negatively related to off-flavor, while α-terpineol was inversely related
to fresh flavor (Golaszewski et al., 1998).
Raspberry (Rubus idaeus, ursinus): The main sugars in raspberry are sucrose, glucose and fructose
with citric as the major organic acid (Robbins and Fellman, 1993). At least 200 volatile compounds have
been identified in raspberry (Honkanen and Hirvi, 1990;, Dourtoglou et al., 1995). Impact flavor
compounds for raspberry are 1-(p-hydroxyphenyl)-3-butanone, cis-3-hexenol, α- and β-ionones, α-irone
and mesifurane. Other abundant volatiles include geraniol, nerol, and linalool among others (Paterson et al.,
1993). The “raspberry ketone” or character impact volatile for raspberry is
4-(4-hydroxyphenyl)-butan-2-one (Larsen and Poll, 1990). It had the lowest threshold (therefore, having
the largest contribution to flavor) followed by α-ionone, β-ionone, geraniol, linalool, and benzyl alcohol.
Furaneol, linalool, and ethyl hexanoate were important general aroma compounds while ethyl butanoate,
methyl butanoate, γ-decalactone and 2-heptanone were important cultivar-specific compounds (Larsen et
al., 1992). The most potent flavor compounds identified using a retronasal aroma simulator in raspberries
were β-damascenone, diacetyl, 1-hexen-3-one, 1-nonen-3-one, 1-octen-3-one, and cis-3-hexenal (Roberts
and Acree, 1996).
Blackberry (Rubus laciniata): Fresh blackberry fruit contain 245 aroma compounds (Georgilopoulos
and Gallois, 1987). The most abundant were heptan-2-ol, para-cymen-8-ol, heptan-2-one, hexanol,
α-terpineol, pulegone, octanol, isoborneol, mytenol, 4-terpineol, carvone, elemincine, and nonanol.
Although heptan-2-ol is an important flavor compound with an intense fruit taste with herbaceous nuances,
no single volatile was identified as blackberry-like (Marton and MacLeod, 1990). Some compounds in
blackberry fruit and leaves are glycosidically bound such as benzyl alcohol, benzoic acid,
3-hydroxy-7,8-dihydro-β-ionol, and cis-3-hexenol among others (Humpf and Schreier, 1991).
Blueberry (genus Vaccinium): Blueberries have glucose and fructose as their major soluble sugars and
citric, malic and quinic acids (Eck, 1986). The odor impact compounds for high-bush blueberry (Vaccinium
myrtillus) are trans-2-hexenal, trans-2-hexenol, and linalool, but also include geraniol, citronellol,
hydroxycitronellol, farnesol and farnesyl acetate. Most volatiles are present below their threshold
concentrations, but hydroxycitronellol was described by sensory panelists as blueberry-like. Rabbit-eye
blueberries (V. ashei) have a different aroma than high-bush. Some aroma volatiles unique to rabbit-eye
blueberries include 1-penten-2-one, γ-terpinene, carveol, acetone, cis-caran-3-ol, ecineralone, α-cedrene,
sabinol, geranyl formate, linalyl acetate, undecan-2-one, tridecan-2-one, ethyl acetate, ethyl tetradecanoate,
dimethyl octanedioate, toluene, p-cymene, and β-ionone among others (Honkanen and Hirvi, 1990).
Grape (genus Vitis): Glucose and fructose are the predominant sugars in grapes, while tartaric and
malic acids account for 90% of the TA (Kanellis and Roubelakis-Angelakis, 1993). Grapes show an
increase in free and glycosylated aroma compounds at the end of ripening, after sugar accumulation has
slowed (Coombe and McCarthy, 1997). This process is different from that of other berries and has been
termed “engusting.” The volatiles in wine grapes are the most complex and are classified into five groups,
of which the first four have glycosylated forms: monoterpene (abundant in “floral” grapes), norisoprenoid,
benzenoid, aliphatic and methoxypyrazine. The accumulation of flavor volatiles occurs late in the berry
ripening cycle, well after accumulation of sugar as observed in Muscat berries (Park et al., 1991). Different
varieties have distinctive aroma character. For example, Muscat odor is mainly composed of monoterpenes
such as linalool and geraniol (Webb, 1981; Kanellis and Roubelakis-Angelakis, 1993). Carbernet
Sauvignon, a V. vinifera cultivar, contains methoxyisobutylpyrazine, which has a strong, green bell
pepper-like aroma (Webb, 1981). Benzyl and 2-phenylethyl alcohols, ethers, aldehydes, and hydrocarbons
also contribute to aroma. American grapes (V. labruscana and V. rotundifolia) are not suitable for wine
production because they possess what has been termed “foxy” and candy-like odors due to compounds like
methyl anthranylate, aminoacetophenone, furaneol and methyl furaneol. ß-Phenylethanol, with its
rose-like odor, was found to be important for muscadine (V. rotundifolia) aroma ( Flora and Nakayama,
1981). The V. vinifera grapes exhibit a mild aroma that is more desirable for wine production (Shure and
Acree, 1995).

Banana (genus Musa)

Sucrose is the predominant sugar in banana initially, but as ripening proceeds, glucose and fructose
accumulate. Malic, citric and oxalic acids are the predominant organic acids with the astringent taste of
unripe bananas being attributed in part to oxalate levels (Seymour, 1993). Characteristic aroma of bananas
arises from a complex mixture of compounds including short-chain fatty acids such as acetates, butanoates,
and 3-methylbutyl esters. Recently nonvolatile glycoside precursors were shown to release glycosidically
bound volatiles from banana pulp by ß-glucosidase, including decan-1-ol, 2-phenylethanol,
3-oxy-pentanoic acid, 3-methylbutanoic acid and benzoic acid (Perez et al., 1997). Esters account for about
70% of the volatile compounds and acetates and butyrates predominate (Seymour, 1993). 3-Methylbutyl
acetate, however, is considered to dominate banana flavor as the key odor-impact volatile (Berger, 1991)
along with butanoate and 3-methylbutanoate (Engel et al., 1990). Unusual phenol derivatives, eugenol,
5-methoxyeugenol, eugenol-methylether, and elemicin contribute background notes for the full-bodied
mellow aroma of ripe bananas (Engle et al., 1990).

Citrus
Sweet orange (Citrus sinensis): The major sugar in most citrus types is sucrose, with varying levels of
glucose and fructose. The major acid is citrate. Typical orange aroma is attributed to alcohols, aldehydes,
esters, hydrocarbons, ketones and other components of which over 200 have been identified. Of these,
esters and aldehydes are the primary contributors followed by alcohols, ketones and hydrocarbons
(Bruemmer, 1975). There is no single impact compound for orange. However, octanal, decanal, nonanal,
dodecanal, ethylbutyrate, and limonene are likely contributors to flavor (Shaw and Wilson, 1980; Shaw,
1991).
Tangerine (Citrus reticulata): Analysis of tangerine essence revealed 34 volatile compounds that were
odor contributors. However, no one compound was found to have a characteristic tangerine odor
(Moshonas and Shaw, 1972). Later studies suggested that the compounds thymol and
methyl-N-methylanthranilate (dimethylanthranilate) are odor impact compounds for this fruit, but that they
are modified by the presence of monoterpene hydrocarbons. Nevertheless, dimethyl anthranilate is the most
potent flavor component (Shaw and Wilson, 1980).
Grapefruit (Citrus paradisi): At least 126 volatile components have been identified in grapefruit
(Demole et al., 1982). Nootkatone and 1-p-menthene-8-thiol may be key aroma impact compounds for
grapefruit (Demole et al., 1982) although aldehydes and esters are also important (Shaw and Wilson, 1980).

Mango (Mangifera indica)

The major sugars in mango are glucose, fructose and sucrose, with sucrose predominating. The major acids
are citric, malic, and sometimes tartaric at 0.1 to 0.4 % TA (Nairain, et al., 1997; Baldwin et al., 1999b,) and
10 to 16 SSC (Baldwin et al., 1999b). Mango varieties differ in amount and type of volatile compounds
present (over 150 compounds identified), often depending on area of production. Asian mangoes have more
oxygenated volatile compounds such as esters, furanones, and lactones, giving some varieties pineapple- or
peach-like aromas (Narain et al., 1997), while Western mangoes that are hybrids of Asian stock have higher
levels of certain hydrocarbons such as 3-carene (MacLeod and de Troconis, 1982; Wilson et al., 1986;
Narain et al., 1997).

Pineapple (Ananas comosus)

Besides banana and possibly mango, pineapple is the most popular fruit from the tropics. SSC can range
from 11 to 17° Brix, and the major sugars are glucose, fructose and sucrose, with sucrose predominating
(Salunkhe and Desai, 1984; Shukor et al., 1998). The major acids are citrate and malate with about 0.1 to
0.6% titratable acidity (Salunkhe and Desai, 1984; Shukor et al., 1998 ). Over 120 volatiles have been
identified in green and ripened pineapples with esters dominating at over 80% of the total volatiles (Shukor
et al., 1998). Contributing aroma volatiles, based on odor thresholds show that pineapple aroma is also
dominated by esters such as ethyl 2-methylbutanoate, ethyl acetate, ethyl hexanoate, ethyl butanoate,
methyl heptanoate and others.

Melons (Cucumis melo)

Sucrose is the principal sugar in most melon types, although high levels of fructose may be present in some
watermelon cultivars. Melons contain citrate and malate, or only malate in watermelon (Seymour and
McGlasson, 1993). Ethyl 2-methylbutanoate and methyl-2-methylpropanoate are among the most
significant contributors to flavor of muskmelon cv. Makdimon, one of C. mello reticulatus cultivars which
exhibit strong characteristic aromas. Muskmelon and watermelon also have cis-non-6-enal and cis,
cis-nona-3,6,-dien-1-ol, respectively. The former has a strong melon-like aroma while the latter is
reminiscent of watermelon rind. 4-Oxononanal and 2-hydroxy-5-pentyltetrahydrofuran have fruity and
green odors and contribute to watermelon aroma. The volatile cis-non-6-enyl acetate has a pleasant
honeydew melon-like aroma (Engle et al., 1990). Other varieties have ethyl 2-methylpropanoate,
2-methylbutyl acetate, 2-methylpropyl acetate and the thioether esters (Wyllie et al., 1995).

Tomato (Lycopersicon esculentum)

The SSC/TA ratio (De Bruyn et al., 1971), or content of SSC or TA are important for flavor (Stevens et al.,
1977; Jones and Scott, 1984). The major sugars are glucose and fructose in roughly equal amounts, while
citrate and malate are the major organic acids, with citrate predominating (Baldwin et al., 1991a and b;
Hobson and Grierson, 1993). However, over 400 volatile compounds were identified, of which 16 or so
have odor thresholds that would indicate that they contribute to flavor (Buttery, 1993; Buttery and Ling,
1993). Of these, there is no clear odor impact compound. Buttery (1993) suggested that a combination of
cis-3-hexenal, hexanal, 1-penten-3-one, 3-methylbutanal, trans-2-hexenal, 6-methyl-5-hepten-2-one,
methyl salicylate, 2-isobutylthiazole, and β-ionone at the appropriate concentrations produces the aroma of
a fresh ripe tomato. Of these, cis-3-hexenal and β-ionone have the highest odor units, and 2-isobutylthiazole
is unique to tomato. Furaneol has an odor threshold indicating it may contribute to flavor (Buttery et al.,
1995). Volatile production occurs at the same time ethylene increases and carotenoid synthesis/chlorophyll
breakdown occur (Baldwin et al., 1991a). Enzymes important in volatile synthesis from lipids include
lipoxygenase, hydroperoxide lyase (hydroperoxy cleavage) and alcohol dehydrogenase (Galliard, et al.,
1977; Riley et al., 1996). Amino acid precursors include alanine, isoleucine, leucine, phenylalanine and
valine (Buttery and Ling, 1993). Glycosides are also precursors to some volatiles (Krammer et al., 1994).
Furaneol is also reported to be important (seems it is found in every fruit). Genetically engineered fruit with
down- or up-regulated alcohol dehydrogenase expression exhibited altered levels of some related volatiles
(Speirs et al., 1998).

Conclusion: Flavor of fruits and vegetables is an important aspect of quality. Although difficult to define,
qualify, and quantify, this elusive and complex trait is important to consumers and deserves more attention
from both researchers and industry. Flavor quality of fresh and processed fruit and vegetable products will
be an important factor in an increasingly competitive global market. Flavor maintenance becomes a
challenge to maintain as shelf life and marketing distances increase due to new storage, handling and
transport technologies. However, despite these issues, the bottom line for flavor quality is still genetic.
Breeders need more information and analytical tools in order to select for flavor quality. Use of wild
material may be necessary in breeding programs to regain flavor characteristics that have been lost from
some commodities. Use of molecular markers that relate to flavor may help identify important enzymes in
flavor pathways. The effect of harvest maturity on flavor quality needs to be determined for each
commodity. With the current focus on flavor quality and current advances in flavor chemistry, sensory
techniques and molecular biology, there are many opportunities to further efforts on behalf of flavor quality
in fresh produce.

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Table 1. Odor descriptors for tomato aroma compounds in deionized water, ethanol/methanol/deionized
water mix and deodorized tomato homogenate (Tandon et al., 2000).

Aroma compound Deionized water EtOH/MeOH/water Tomato

homogenate

hexanal grassy/green rancid/stale oil stale/green/grassy

trans -2-hexenal floral/grass/apple fruity/almond/vine stale/green/vine

cis-3-hexenol leafy/cut grass freshcut grass green/celery

hexanol mint/grass alcohol glue/oil

6-methyl-5-hepten-2-one raw greens/nutty alcohol/paint sweet/floral

cis-3-hexenal grass/tomato-like alcohol/paint tomato/citrus

2-isobutylthiazole fermented/plastic alcohol/tomato-like pungent/bitter

2-pentenal vine/organic solvent acetone/medicine stale/oil

acetone glue/alcohol alcohol/nutty/spoilt green

β-ionone sweet/perfume-like sweet sweet/floral

geranylacetone sweet/paint/sharp sweet/floral/leafy sweet/citrus/ester

3-methylbutanol earthy/watermelon rind glue/mint/cinnamon sweet/fresh

phenylethanol floral/roses alcohol alcohol/nutty

3-methylbutanal bug spray/alcohol fruity/green/leafy stale/rotten

1-penten-3-one glue/oil/pungent nutty/glue/alcohol fresh/sweet

ethanol earthy/stale pungent/rancid

methanol earthy/stale
Table 2. Some important or abundant flavor compounds in selected fruits and vegetables.

Fruit Major Major acids Important aroma compounds

sugars
Apple sucrose malic β-damascenone
glucose citric butyl hexanoate
Fellman et al., 1993 fructose isoamyl hexanoate
Honkanen and Hirvi, 1990 hexyl hexanoate
Knee, 1993 ethyl butanoate
Mattheis et al., 1995 propyl butanoate
Young et al., 1996 hexyl butanoate
butylacetate
2-ethyl-1-butyl acetate
ethyl acetate
butanol
Peach sucrose malic benzaldehyde
glucose citric benzyl alcohol
Brady, 1993 fructose nonanol
Crouzet et al., 1990 sorbitol linalool
Do et al., 1969 ethyl hexanoate
3-methylbutanoate
α-terpineol
γ-hexalactone
δ-decalactone
γ-undecalactone
δ-undecalactone
γ-dodecalactone
δ-dodecalactone
α-pyrone
6-pentyl-α-pyrone
Strawberry sucrose, citric hexanal
glucose, cis-3-hexanal
Golaszewski et al., 1998 fructose trans-2-hexanal
Honkanen et al., 1980 furaneol
Manning, 1993 mesifuran
Perez et al.,1999 ethyl hexanoate
Roscher et al., 1997 ethyl butanoate
Zabetakis and Holden, 1997 methyl butanoate
ethyl-2-methyl propanoate
Raspberry sucrose citric H-(4-hydroxyphenyl-butan-2-one)
glucose (raspberry ketone)
Dourtoglou et al., 1995 fructose α-ionone
Honkanen and Hirvi, 1990 β-ionone
Larsen and Poll, 1990; Larsen et geraniollinalool
al., 1992 benzyl alcohol
Paterson et al., 1993 ethyl hexanoate
Robbins and Fellman, 1993 ethyl butanoate
Roberts and Acree, 1996 methyl butanoate
γ-decalactone
2-heptanone
cis-3-hexanal
β-damascenone
Grape glucose tartaric methyl anthranilate
Concord fructose malic 0-aminoacetophenone
(Vitis labruscana) furaneol
methyl furaneol
β-damascenone

Muscadine β-phenylethanol
V. rotundifolia butyl alcohol
hexyl alcohol
(References for all grape types) hexanal
Coombe and McCarthy, 1997 trans-2-hexenal
Flora and Nakayama, 1981 isoamyl alcohol
Kanellis and Roubelakis- acetaldehyde
Angelakis, 1993 isobutyraldehyde
Park et al.,1991 ethyl acetate
Shure and Acree, 1995 ethyl propionate
Webb, 1981 butyl acetate
propyl acetate
2-methylbutanol

Muscat varieties linalool

(V. vinifera) geraniol
methoxyisobutylpyrazine
Banana sucrose malic decan-1-ol
glucose citric 2-phenylethanol
Berger, 1991 fructose oxalic 3-oxy-pentanoic acid
Engel et al., 1990 3-methylbutanoic acid
Perez et al., 1997 3-methylbutyl acetate
Seymour, 1993 butanoate
3-methylbutanoate
eugenol
5-methoxyeugenol
eugenol-methylether
elemicin
Sweet Orange sucrose citric geranial
Bruemmer 1975 glucose neral acetaldehyde
Shaw 1991 fructose decanal
Shaw and Wilson 1980 octanal
nonanal
ethyl acetate
ethyl propionate
ethyl butanoate
methyl butanoate
 ethyl-2-methyl butanoate
ethyl-3-hydroxy hexanoate
linalool
α-terpineol
limonene
myrcene
α-pinene
valencene
Tangerine sucrose citric acetaldehyde
glucose decanal
Moshonas and Shaw, 1972 fructose octanal
Shaw and Wilson, 1980 dimethyl anthranilate
thymol
α-sinensal
γ-terpinene
β-pinene
Grapefruit sucrose citric acetaldehyde
glucose decanal
Demole et al., 1982 fructose ethyl acetate
Shaw and Wilson, 1980 methyl butanoate
ethyl butanoate
1-p-menthene-8-thiol
nootkatone
limonene
naringin
Mango sucrose citric ethyl butanoate
glucose malic ethyl-2-butanoate
Baldwin et al., 1999 fructose hexanal
MacLeod and de Troconis, 1982 cis-3-hexanal
Nairain et al., 1997 trans-2-hexanal
Wilson et al., 1986 γ-octalactone
γ-dodecalactone
furaneol
α-pinene
β-pinene
3-carene
myrcene
limonene
p-cymene
terpinolene
α-Copaene
caryophyllene
Melon: sucrose malic ethylbutyrate
Cantaloupe fructose citric ethyl-2-methyl butyrate
Honeydew watermelon - ethyl butyrate
Watermelon malic only ethyl hexanoate
hexyl acetate
Engle et al., 1990 3-methylbutyl acetate
Seymour and McGlasson, 1993 benzyl acetate
Wyllie et al., 1995 cis-6-nonenyl acetate
trans-6-nonenol
cis, cis -3,6-nonadienol
cis-6-nonenal
4-oxononanal
2-hydroxy-5-pentyltetra-hydrofura
n
cis-non-6-enyl acetate
methyl acetate
ethyl acetate
isopropyl acetate
ethyl propanoate
ethyl isobutanoate
propyl acetate
butyl acetate
methyl-2-methylbutanoate
ethyl butanoate
2-methylpropanoate
2-methylbutyl acetate
2-methylpropyl acetate
methyl (methylthio) acetate
ethyl (methylthio) acetate
ethyl (methylthio)propanoate
Tomato glucose citric hexanal
fructose malic trans-2-hexenal
Baldwin et al.,1991a and 1991b cis-3-hexenal
Buttery, 1993 cis-3-hexenol
Buttery and Ling, 1993 β-ionone
Buttery et al., 1995, 1989 β-damascenone
De Bruyn et al., 1971 1-penten-3-one
Hobson and Grierson, 1993 3-methylbutanal
3-methylbutanol
2-isobutylthiazole1-nitro-phenyl-
ethane
trans-2-heptenal
phenylacetaldehyde
6-methyl-5-hepten-2-one
methyl salicylate
geranylacetone